The British warblers : a history with problems of their lives

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1-6 AQUATIC -W. 7-12 SEDGE-W.

PUBLISHED BY

WARBLERS,

139-18 REED -W.
25-34 GREAT REED -WARBLER.

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19-24 MA

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FEBRUARY, 1907.

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A HISTORY WITH PROBLEMS
Fae mes OF

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“He ELIOT HOWARD, F.Z.S., M.B.O.U.
ieee “InbusrrarEp BY HeNRIK GRONVOLD :

porta ! eae London
ee ee eR PORTER
: 7, PRINCES STREET, CAVENDISH SQUARE, W.

Price 21s. net.

SMITHSON Tap

JAN 23 1986
Ss LIBRARIES”

ITISH WARBLERS,

- CONTENTS. )

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; : - Sedge- Warbler peer years Tot aie Be oY Noe eae
ne < - Grasshopper-Warbler eotes, ee at re es yy, 124

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PLATES.

eet oe Eggs of British Warblers. Plate I. (coloured).
ay =o pees Sedge- Warbler ad. ¢ (coloured).
| — Pits ee 4 » & (Photogravure).
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tee Grasshopper-Warbler. ,, ¢ (coloured).
om es » & (Photogravure).
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=a ; % Nestlings ,,
: re TWO MAPS

- Showing approximate Geographical Distribution of Grasshopper-
and Savi’s Warblers, during Summer and Winter.

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ADULT MAL

23

SEDGE-WARBLER.

Sylvia phragmitis, Meyer, British Birds, folio Bd., vol. i. (coloured
plate figuring adult and egs) [1835-43].

Salicaria phragmitis, Hewitson, British Oology, 1st Ed., vol. i,
pl. 70, fig. 2 (ege) [1836]; 2d., Hogs of British Birds, 2nd
Ed., vol. i, p. 87, pl. 25, fig. 2 (egg), 1846; zd. 2d., 8rd Ed.,
vol. i, pp. 117-118, pl. 31, fig. 3 (egg), 1856; Booth, Rough
Notes, vol. ii, pp. 43-44, 1888.

Calamoherpe phragmitis, Macgillivray, British Birds, vol. ii,
pp. 390-394 (woodcut of head), 1839.

Calamodyta phragmitis, Gould, Birds of Great Britain, vol. ii,
2 pp, pl. 75 (coloured figures of adult male and female), 1862.

Acrocephalus schoenobenus, Yarrell, British Birds, 4th Ed.,
vol. i, edited by Newton, pp. 376-379 (woodcut), 1878 ;
Dresser, Birds of Hurope, vol. ii, pp. 597-601, pl. 90, fig.
2 (coloured figure of adult male), 1876.

Acrocephalus phragmitis, Seebohm, British Birds, vol. i, pp. 352-
306, pl. 10, fig. 17 (egg), 1883; Lilford, Coloured Figures,
vol. ili, p. 40, pl. 20 (coloured figure of adult male), 1886 ;
Saunders, Manual of British Birds, 2nd Hd., pp. 85-86
(woodcut), 1898.

Arabian, Fissew; Croatian, Vodarisa rogocara; Czechisch, Mysak ;
Danish, Swsanger; Dutch, Rietzanger; Finnish, Ruohokerttu; French,
Bec-fin phragmite ; German, Schilf-Rohrsdinger ; Hungarian, Foltos sitke ;
Italian, Forapagle; Maltese, Violin; Norwegian, Sivsanger; Polish,
Trzcumak rokit-miczka; Russian, Kamyschefka kamyschewaja; Spanish,
Buscarla; Swedish, Sdfsangare.

DESCRIPTION OF THE PLUMAGE.

Adults in Spring. — The sexes are alike, the male,
perhaps, being a trifle brighter and rather larger, but on
the other hand a bright female may surpass a dull-coloured
male. The upper parts are of an umber brown tint shad-
ing into an almost unspotted rusty brown on the rump and
upper tail-coverts.. There is a conspicuous buffish white
superciliary stripe and above that an equally conspicuous
blackish ‘stripe; the feathers in the middle part of the crown

1

BRITISH WARBLERS

have dark centres forming three longitudinal stripes, the
lores are blackish brown, cheeks brown, and the upper parts
of the ear coverts slightly more dusky, these latter forming
with the lores a darkish stripe through the eye. The
hinder part of the neck is almost unspotted, whereas on
the back and shoulders each feather has a blackish brown
centre forming about six stripes on the back. The wing-
coverts are blackish brown with umber brown edges to each
feather, the flight-feathers are brown with slightly lighter
edges, the innermost secondaries having blackish brown
centres and light umber brown edges, the primaries having
narrow whitish brown tips. The upper part of the tail is dark
umber brown with lighter edges of the same tint, both colours
getting lighter towards the outermost rectrices. The throat
is whitish, the crop and sides of the neck buff shading into
umber buff on the flanks and under tail-coverts. Abdomen
whitish, and the under part of the tail greyish brown with
a wash of lavender grey.

Immature.—The colour of the young is much like that
of the adults, but richer. The crown is blackish, each feather
narrowly edged with olive buff, the combined effect being,
when the feathers are in perfect order, that of six fairly broad
blackish stripes divided by narrow olive buff lines. There is
a conspicuous light buff superciliary stripe, the lores are
blackish, uppermost ear-coverts dusky, the combined effect
being a dark streak through the eye; on the nape the blackish
centre and olive buff edges of the feathers are of equal
strength ; on the back and shoulders the blackish centres
are most conspicuous, forming about six fairly well defined
longitudinal stripes; on the rump and upper tail-coverts the
olive buff becomes slightly rusty or richer olive buff and the
dark centres to the feathers are less conspicuous. The upper
part of the tail is brownish slate, each feather edged with
olive buff slightly lighter on the outermost ones, the shafts
being dark lavender brown. The wings are darkish brown
slate, each feather boldly edged with olive buff, narrower on

2

SEDGE-WARBLER

the small coverts, and forming a sort of dark band across the
wing, almost hiding the dark centres on the upper wing,
making this part look uniform in colour. The sides of the
face and neck are rather rich olive-buff with faint dark centres
to the feathers, and there is a moustache stripe, though not
very conspicuous. The throat is pure whitish buff; the crop
and flanks are buff, with or without a faint olive wash, the
former being furnished with brownish grey spots. The
abdomen is whitish, blending into a pure buff on the under
tail-coverts; under surface of the tail and wings are lavender
grey narrowly edged with light buff and the feather shafts
are white. The upper mandible is dark horn lavender, lower
mandible light lavender flesh, darker towards the tip, and
the corner and inside of the mouth orange yellow. Irides
dark brown, the small feathers above are light buff, those
below white. The eyelid is dark lavender flesh. Tibia olive
buff. ‘Tarsus and toes fairly light greenish lead colour; goles
yellowish olive, and the claws olive grey.

GEOGRAPHICAL DISTRIBUTION.

It is so generally distributed from the middle of April to
September throughout the British Isles that it is not necessary
to mention any particular locality: we find, however, that as
we proceed further north it is rather more local, and does not
apparently reach the Shetlands, although in the Orkneys it is
by no means uncommon. In the north-west of Skye it is
rare, but in the islands further south, Islay and Mull, more
numerous. In Ireland it breeds in every county, and is even
found on Achil Island.

Crossing the Hnglish Channel and commencing in the
south-west, we find it rare in the south of Spain and in
Portugal, but rather more numerous further north, especially
near Aroza Bay. In the Pyrenees it is common, also in
suitable places throughout France, Belgium, and the Nether-

3

BRITISH WARBLERS

lands, especially, in the latter countries, along the banks of
the Rivers Scheldt and Meuse. I did not find it as common
as I should have expected in the Island of Texel: it passes
over Heligoland in great numbers.

Continuing northwards, we find it generally distributed

throughout Denmark and breeding in suitable localities. In
the south of Sweden it is common, then becomes rarer as far
as the south of Norrland, but north of this it apparently does
not occur: it is unknown in Gothland. In the south of
Norway it is rare, but has been observed in the Jaderen
district and up to Laurgaard; north of this it disappears
until we reach Dynnisé in the Helgeland district, where it
becomes more common again and reaches as far north as
Finmark.
_ Returning to Central Europe, we find it generally dis-
tributed over Germany and especially common in the marshy
parts of Mecklenburg, Holstein and Westphalia; also in
Hessen and along the banks of the Rivers Moselle and Elbe;
rare, however, in Sachsen-Altenburg, but very common in
Silesia. In Switzerland it is fairly numerous, especially in
the low-lying country and valleys round Geneva and Lake
Constance, but in the central and northern parts it is rarer,
although inhabiting some of the lower sub-alpine valleys,
especially Hasli. In Italy it is common in all the marshy
parts, but not so numerous in Sardinia, and rare in Corsica.
Kastwards, we find it again common in the Austro-Hungarian
Monarchy, especially in the central plains along the banks
of the Danube. Whether it breeds in Montenegro is doubtful,
but immense numbers pass on migration. On the east side
of the Balkan Peninsula we find it common in the reed-
beds of Varna and Pravadia in Bulgaria, but further inland
less numerous.

In Greece it is a bird of passage only, appearing on
migration in large numbers from the end of March to the
end of May. It may, however, have been overlooked as a

a.

SEDGE-WARBLER

breeding species, since there are still extensive unexplored
Swamps in the country.

The Russian Empire is a large breeding ground. In
Poland it is the most common of all the reed-warblers, and
it is also numerous in the Baltic provinces. Crossing the
Gulf of Finland, we find it common in suitable places in the
provinces of Nyland and Tevastehus, in the neighbourhood
of Abo and Bjérneborg, near Lake Ule&, and on the Island
of Carlé6; and still further north breeding in the Kola
Peninsula, but rare on the shores of Lake Enara. It occurs
north-east of Lake Ladoga, in the province of Olonetz, on the
banks of the River Svir and near Lake Onega. In the vicinity
of Archangel it is numerous, and it also occurs in the valleys
of the Petchora, Ob and Yenesei.

In the provinces of Pskov, St. Petersburg, Smolensk,
Jaroslav and Kasan it is common, scarce in Novgorod, Tver
and Riazan, and local in Moscow and Tula.

In the central valley of the Volga it is numerous, Ago in
places in the Ural districts, but in the province of Orenburgh
it is local, and rare in the Kirghiz Steppes, and again we find
it numerous in Astrakhan and the delta of the Volga; and
continuing south we find it occurring in Stavropol, Terek and
Trans-Caucasia, and crossing the Caspian Sea inhabiting the
Mangishlak Peninsula, Ust Urt plateau, and the valley of
Amu Daria.

In Turkestan it is migratory and breeds, and we trace it
eastward to the Altai Mountains.

In Palestine itis common in suitable places, also in many
parts of Africa, breeding in Algeria, Tripoli, Tunis and Egypt.

The winter home is principally Central and Southern
Africa, but it probably also remains in some of the countries
bordering on the Mediterranean, since it has been found at
this season in Dalmatia.

In Lower Egypt it is common, and has been found in
Somaliland and on Lake Haramaia, and we also trace it to
German Hast Africa and as far south as Damaraland.

3)

BRITISH WARBLERS

LIFE-HISTORY.

Arriving in this country about April 25th, these birds
choose many varied spots for their home; osier-beds, where the
various species of sedge grow in abundance, they favour most,
although at times they will choose a thick and tangled hedge-
row, and again, and this very frequently, they may be found
amongst the dense masses of the Arwndo phragmutis, but in
the drier portions. The arrival of the males, and perhaps of
the females also, is rather irregular; that is to say, a few will
appear one morning, then there will be a pause, and a few
days later another batch will arrive, and so on until the
migration of the males ceases, about a fortnight after the
arrival of the first individual.

Their arrival can soon be detected by their babbling sone,
an energetic but unmusical strain, and where first heard there
they will probably be found to breed, for they are by nature
most home-loving individuals.

Water or swampy ground seems to be a necessity for
them, for they are rarely to be found breeding in dry places.
I have occasionally come across them in small dry coppices,
but never very far away from water of some description.
They inhabit more particularly wet osier-beds, where the
different sedges, the Carex ovalis, Carex acuta and Carex
ryparva grow abundantly, but they seem to prefer them when
rather drier, that is to say, where the willows are young
and thick, and where the Juncus effusus, Spirea ulmaria
(meadow sweet) and different species of Hpilobiwm (willow
herb) grow in tangled masses. Along the banks of rivers and
streams they are often common, and are abundant on pools
and lakes, where aquatic plants, such as Scirpus lacustris
Typha tatyfolia (bullrush) and the tall reeds (Arwndo phragmitis)
grow. Where these reeds grow to a great height, as they do
in Holland and Hungary, and where the bottom is a dry mass
of roots raised above the water, there I have found them
exceedingly common.

WARBLER,

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SEDGE-WARBLER

The males, when they first arrive, select a certain spot of
not many square yards in extent, with a tall bush or willow
conveniently situated; and, moreover, they not only choose a
tree but some particular branch on that tree, and this, until
the young are hatched, forms their headquarters. Before the
female arrives, and also, but not as much, when paired, they
sit on this particular branch and pour out their song,
frequently accompanying it with a pretty erial flight, rising
almost perpendicularly for a short distance in the air, turning
very quickly and returning, with wings and tail outspread,
to the branch. After a short rest they either start their song
afresh, begin to preen their feathers, or go in search of food,
wandering over the branches of the various willows, examining
the under parts of the leaves for flies of the genus Chironomus,
and the twigs for aphides, working lower and lower down
through the branches and thick herbage till they reach the
ground, and here for a time they make their way in search of
aquatic insects and small spiders, only shortly to return to
their favourite branch, singing as they fly. Thus they while
away the days till the females arrive. The ground imme-
diately surrounding the tree they have chosen as their
headquarters they look upon as particularly their own, and
when two or three have chosen positions close to one another,
there they will be found to search for their food in certain
well-defined directions, being most careful not to poach on
one another’s preserves, and consequently most jealous of any
intrusion on the part of their neighbours, and not, indeed,
only of- their neighbours, but also—and this I have seen so
frequently as to place it beyond the bounds of chance—
of any other warbler, chasing them determinedly away,
while frequently—I have seen them pursuing Thrushes and
Hedge-Sparrows—ignoring the presence of other species so
far as to allow them to nest in their favourite tree. Such
powers of apparent discrimination seem very wonderful,
yet we must not forget our total ignorance of all things
pertaining to the sub-conscious state of animal life, and

7

BRITISH WARBLERS

consequently of the bearing it may have on the unknown
connection between different individuals, or classes of
individuals, which, if known, would go far to explain such
incidents, and perhaps enable us to form a truer definition of
species than hitherto possible.

The courtship commences directly the females arrive, about
ten days or so after the males; and as the females at this
period persist in skulking in the bottom of the thickest rushes
and undergrowth, it is most difficult to see what actually takes
place; but the glimpses I have occasionally obtained tend to
make me look upon the males when in love as rather sober-
minded individuals ; although the females for their part insist
upon very close attention, and a somewhat servile attitude
on the part of their suitors. A male pursues the female with
drooping wings and erected head-feathers, uttering a rather
harsh call-note; sometimes when quietly following he picks
up and carries a dead leaf. If another male approaches too
closely he pursues him with quick, vigorous flight. The
female all the time keeps up her gentle call-note, which
_ becomes more vigorous if her suitor, while pursuing another
male, is forgetful and leaves her too long alone. Sometimes
the two play together, flying at one another, the male
scolding; and again they will sit close beside one another,
an uninteresting couple, except when the male runs up and
down the branch sideways, as he frequently does, with
drooping wings and tail.

The nest, placed low down amongst the thick tangled
undergrowth in the fork of a willow, or on the low branches of
a bramble, especially when the latter entwines amongst a thick
growth of Juncus effusus, is built up as follows: The founda-
tion is dead grass of various kinds mixed with small pieces of
dead thistle; on this is a thick layer of the fluffy seeds of the
various species of Salix, bound together with fine dried grass,
the lining being usually of the latter substance only. The
female does most of the building, flying backwards and for-
wards to the Salia nigricans, carrying billfuls of the seeds,

8

SEDGH-WARBLER

followed closely by the male, who never makes the slightest
effort to help her, but, whether she be on the willow tugging
at the seeds or actually building, sits close beside her, and in
the same manner flies within afew feet of her on her journeys
backwards and forwards.

The young are hatched as a rule about the middle of June,
but the date varies considerably ; I have found them as early
as the first week of that month, and first broods as late as the
first week in July. When the young are hatched the parent
birds will allow you, if well concealed, to watch their domestic
arrangements very Closely. Peering through the sedges
within a few feet of the nest, I have frequently watched them
feeding their young, and cleaning their nest. The female at
this time is less suspicious than the male. She it is who
seems to be aware of the necessity of a constant supply of food
to the nestlings; the male in this respect not working nearly
so hard, and when suspicious only occasionally, and that very
hurriedly, bringing a small billful of insects: the greater part
of his time appears to be spent in keeping guard. The female,
on the other hand, feeds her brood continually, every few
minutes coming to the nest with a large billful of insects,
which she distributes amongst some of the young. When
startled she will sing a few notes of the song of the male, but
will on no account drop the food she may be holding at the
time. I have startled her in many ways, even going so far as
to touch her with a stick, in order to make her drop the insects
that I might be able to see to what species they belonged, but
in no case have I been successful. The feces, enclosed in a
membranous sac, is carried away by the female each time
alter bringing food, and is dropped twenty or thirty yards from
the nest ; occasionally, but not as frequently as amongst other
species, 1b is swallowed by her. The young leave the nest
when about ten days old, and until able to fly keep well hidden
amongst the undergrowth, occasionally uttering their very
small call-note, which is answered by the parent birds, and
enables them to keep in touch with their scattered offspring.

9

BRITISH WARBLERS

When fully grown the young ones are soon taught to find
food for themselves. They follow their parents amongst the
bushes and undergrowth, each being fed in turn, but occa-
sionally taking the trouble to pick off a few aphides for them-
selves, giving one the impression that they prefer to be lazy as
long as possible. Their difficulty no doubt lies in their want
of experience, and consequent lack of power to distinguish
between what is food and what is not; thus, even when they
have left their parents, I have seen them in doubt trying to
swallow the ragged ends of a broken twig. They are very
playful at this age, their games sometimes taking the form of
a tilting match. Three take part, two sit on convenient twigs
facing one another, and the third from his central position
might almost be called an umpire. Numbers one and two
then lower their heads, each in anticipation of the other
moving; one of them, call him number one, then springs into
the air, and darts at number two; number two dodges and
occupies the position vacated by number one, each of them
then face round ready to continue the fray, the change of
positions becoming quite rapid.

Although I have never found a second brood, yet I think
it probable that two are sometimes reared, for this reason,
that some of the males in July indulge in a second court-
ship. When a female is present the male will spread out
his wings and tail, and walk sideways up and down a branch,
the female for her part also spreading out her wings and tail,
but not moving up and down ‘the branch. After perform-
ing thus for a short time she will fly off, pursued by the
male, calling to him with her low quiet note if he does
not immediately follow. Sometimes two or three males join
in the pursuit, apparently for the fun of the thing and the
annoyance of the real lover.

It will be seen that this courtship is really identical with
that performed in spring, and is therefore somewhat of a
mystery, the more so as it is not indulged in by all males, for

10

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SEDGE-WARBLER

some at this period, after they have reared their young, begin
to wander, and can be heard singing even in the middle of the
cornfields.

Towards the middle of July, while watching them in osier-
beds, where they happened to be very plentiful, I have noticed
curious commotions, the origin of which I have not been
able to satisfactorily explain. As many as eight or nine
collect together, some adult, some immature, the adults
apparently very angry, scolding vigorously, their feathers
ruffled and their tails spread out; the onlookers also join in
the scolding, the commotion being quite unusual for bird-
life. So absorbed have they sometimes been that I have crept
into the middle of them before they have noticed me, when
they have reluctantly dispersed. 1 once found an empty nest
from which the young may have been taken, but there were
no dead bodies round it to show that a tragedy had occurred ;
and it is generally the case, when a rat or a weasel has
attacked and destroyed the young, that the bodies lie round
or perhaps even remain in the nest, with no sign of ill-usage
on them except two small punctures at the junction of the
head and neck, indicating the marks of the teeth, the blood
having evidently been sucked. Hitherto, I have seen these
scolding parties in July only, but it is quite likely that they
may be of frequent occurrence at other times.

On his arrival in this country the male sings at all hours
of the day, especially in the morning, frequently at night, and
continues until pairing is over and incubation commenced,
when to a great extent he ceases, but recommences after the
young have flown. His song is loud, cheerful and babbling,
sometimes harsh, and perhaps might be described as lacking
‘education. He has also considerable powers of mimicry,
the call-notes and parts of the song of different species
being often introduced. This mimicry will be heard more
frequently when any commotion is going on amongst the
bird-life around, owing, no doubt, to the alarm-notes of the
different species engaged in it being then more pronounced.

I

BRITISH WARBLERS

At such times I have heard them imitate the Whitethroat,
Blackbird and Chaffinch, and at other times the Tree Pipit,
while once I remember hearing one imitating the call-note
of the Partridge perfectly. They can also, when singing, pro-
duce a ventriloquistic effect, but do not do so very frequently.
The ordinary call-note used by both sexes is a rather harsh
crackling note, difficult to describe; the female when mating
has rather a different note to the male, and the young when
fully grown differ again somewhat from both their parents.

The call-note—this is rather a vague term, in reality
meaning very little—is used by both sexes all the summer,
and when listening by the side of a reed-bed on hot summer
afternoons it is frequently the only expression of bird-life one
hears. Thus they keep calling, sometimes with the single
note, sometimes with a treble note quickly uttered in a
descending scale, as they wander along in search of food,
apparently not troubling about an answer to their call, but
uttering it mechanically at varying intervals. Any sudden
noise or disturbance, even the throwing of a stone into the
rushes, is often enough to cause them to sing.

In the latter part of August and the first half of September,
that is to say, in the last few weeks before they leave this
country, they are much quieter, rarely singing, but instead
skulking in the undergrowth, and are therefore difficult to
see. As they move about you can hear them occasionally
calling to one another, but showing themselves very little,
evidently finding plenty of food amongst the stems of the
reeds and rushes.

Climatic conditions do not seem to affect them very much,
but during very wet weather their song is not so frequent
nor so vigorous, neither do they appear to be very happy
themselves, but are rather more inclined to mope and are
considerably less active.

Their food seems to consist entirely of insects, and during
the few months they are with us food of this description is
never lacking. They search for it chiefly low down amongst

12

PUBLISHED BY R.H.PORTER.

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SEDGE-WARBLER.

the roots of the herbage, often hopping on to and along the
ground in swampy places, where small worms and spiders are
abundant, and since they generally inhabit swampy places,
swamp-loving insects form a great portion of their food. It
is common to see the adults hunting the willows and small
alders (Alnus glutinosa), but never the high trees. The young
in July when fully grown are not so particular in this respect,
often searching in the tops of large alders, especially during
the swarming time of the aphides. It is no doubt the pursuit
of food that leads the adults into the middle of cornfields, but
I have not been able to determine what the special food
they are following consists of.

Flies belonging to the Chironomide, of which I have
identified the following, are frequently taken: Chironomus
viridis, Chironomus brevitibialis, and Tanypus choreus.

These insects can be seen, especially in the morning, both
early and late in the season, clustering under the leaves of the
willows, hazels and alders. A slight tap on the branches will
frequently produce incredible numbers, where before few could
be seen.

In June and July they feed, amongst other insects, on the
following :—

GENUS. SPECIES.
Homoptera... ... COrcadula septennotata.
Hemupteron ... ... Macomma ambulans.
Coleoptera... ... Cyphon coarctatus.

Dolichopus wngulatus.
Tiwpule.
ENED Limnobude.
Empide.

The young, especially when still in the nest, are fed largely
on these insects, occasionally also on the larve of certain
moths and butterflies. During all the summer months great
numbers of aphides are devoured, in fact, these insects appear
to constitute the great food supply, not only of this species,
but of many others also. A close inspection of the twigs and

13

BRITISH WARBLERS

leaves of the different species of Salix will reveal these insects
clustering in great numbers, and will at once explain what,
at a distance, looks like the aimless pecking at nothing of
these birds.

When inhabiting swamps where the Arwndo phragmitis
grows, they seem to find, early in the season, a quantity of
food amongst the fluffy seeds at the top of these reeds.

14

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25

GRASSHOPPER-WARBLER.

Sylvia locustella, Meyer, British Birds, folio Ed., vol. i. (coloured
plate of bird and egg) [1835-43].

Salicaria locustella, Hewitson, British Oolog Vy 1st Hd., vol. i, plate
70, fig. 8 (egg) [1836]; zd., Hggs of British Birds, 2nd Ed.,
vol. i, pp. 85-86, pl. 25, fig. 1 (egg), 1846; zd. ad., 3rd Ba.,
vol. i, pp. 112-114, pl. 31, fig. 1, 1856; Booth, Rough Notes,
vol. li, pp. 41-42, 1883.

Sibilatrix locustella, Macgillivray, British Birds, vol. ii, pp.
399-404 (woodcut of head), 1839.

Locustella avicula, Gould, Bids of Great Britain, vol. ii, 2 pp,
pl. 78 (coloured figures of adult male and female), 1866.

Acrocephalus neevius, Yarrell, British Birds, 4th Ed., vol. i, edited
by Newton, pp. 384-388 (woodcut), 1873.

Locustella neevia, Dresser, Birds of Europe, vol. ii, pp. 611-616, pl.
91 (coloured figures of adult male and young bird), 1874;
Inlford, Coloured Figures, vol. iii, p. 44, pl. 22 (coloured
figure of adult male), 1888; Sawnders, Manual of British
Birds, 2nd Ed., pp. 89-90 (woodcut), 1898.

Locustella locustella, Seebohm, British Birds, vol. i, pp. 340-345,
pl. 10, fig. 19 (egg), 1883.

Croatian, Trotenjara kobilicarka ; Czechisch, Rdkosnik zeleny ; Danish,
Busksanger; Dutch, de Sprinkhaan-Rietzanger; French, Bec-fin locus-
telle; German, Heuschrecken-Rohrsdnger ; Hungarian, Réti tiicsékmadar ;
Italian, Forapaghe macchiettato; Norwegian, Greshoppe-Sanger, Busk-
Sanger; Polish, Trziniak swierseceyk; Russian, Swertschok ; Swedish,
Grdshoppsangare.

DESCRIPTION OF THE PLUMAGE.

Adult Male in Spring.
darkish red olive brown, each feather having a blackish brown

iL

BRITISH WARBLERS

centre, and this centre is most conspicuous on the crown
and back, fainter on the long upper tail-coverts, and almost
disappears on the back of the neck. The upper surface of the
tail is of the same hue as the upper parts, but of a slightly
darker brown, narrowly edged with lighter olive brown, and
marked with faint transverse bars—the shaft of the feathers
being dark reddish brown. The lores are olive grey, a super-
ciliary streak, which disappears behind the eye, is a whitish
olive yellow ; cheeks and ear-coverts are darkish olive brown,
with very indistinct light shaft stripes, and the side of the
neck is the same colour as the upper parts but slightly lighter
and unspotted. The wing-feathers are the same colour as the
upper surface of the tail, broadly margined with the same
colour as the back, and slightly reddish at the root of the
outer primaries, forming an indistinct patch, but disappearing
at the tip of the primaries and inner web of the secondaries.
The throat is whitish buff, crop region buffish olive yellow,
centre of breast and abdomen whitish, blending into whitish
buff on the long under tail-coverts, the latter being furnished
with a dark centre, which, however, does not reach to the tip
of the feathers. The under surface of the tail is brownish
grey, suffused with lavender grey. The sides of the breast
and body are of the same colour as the back, but rather
lighter and unspotted, though at the roots of the tail the
feathers have dark but not very conspicuous centres.

Adult Male in Summer.—The upper parts are of an olive
greyish green, very slightly tinged with yellowish umber, each
feather having a dark brown centre, most conspicuous on the’
crown and back, but more indistinct on the nape and upper
tail-coverts. The upper part of the tail is uniform, the olive
umber being stronger, and the shafts a lavender brown. ‘The
wings are a dark greyish brown, but the broad margins of olive
greyish green make the closed wing look almost uniform in
colour; the first and second primary each have a whitish brown
edge. ‘There is an indistinct whitish ochre superciliary stripe
disappearing behind the eye. The lores are greyish, sides of

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GRASSHOPPER-WARBLER

the head and neck olive greyish green. The under parts vary,
the throat being generally whitish and spotted, or unspotted ;
the crop varies from light buff to deep olive buff. The middle
of the abdomen is whitish, often washed with light buff. The
flanks are of a stronger olive brown; the under tail-coverts
vary from light buff to light olive brown, with dark brownish
arrow-headed spots. Under parts of the tail-feathers are
brownish lavender grey with indistinct crossbars and whitish
shafts to the feathers. The upper mandible is dark brown,
the lower horn green, lighter towards the tip ; iris dark brown ;
legs fairly light brownish flesh, the soles having a wash of
olive yellow.

Adult Female in Spring.—The upper parts are of a uniform
lightish olive umber slightly washed with a rusty colour. The
marking on the back and shoulders is the same as in the
male and well pronounced, the dark centres to the feathers
forming on the crown about six narrow stripes, which, though
plain, are not very conspicuous. The back of the neck is
unspotted and the dark centres are very inconspicuous on the
rump and upper tail-coverts, whereas the transverse bars on
the tail-feathers are easily seen. The wing is brownish black
on the innermost secondaries, with a fairly broad edge of the
same colour on the upper parts, though more rusty. The
outer edge of the first large primary is whitish buff. The lores
are dusky, and there is an indistinct superciliary stripe. ‘The
feathers on the eyelid are whitish buff, throat whitish, sides
of the head, neck and body rich gamboge olive, darkening
towards the tail, with faint dark centres to each feather, and
blending into whitish on the centre of the abdomen. The
crop is fawn colour, with small brown spots, and suffused with
a beautiful vinous tint. The under tail-coverts are whitish
buff with dark brown centres, both colours well pronounced.
The under part of the tail is brown, washed with a faint
lavender grey.

Fledglings.—The upper parts are very much like the
adults, only slightly richer in tint and distinctly olive rust

3

BRITISH WARBLERS

colour—not olive grey. There is a distinct, though not
prominent, yellowish superciliary stripe, which soon disappears
behind the eye. The ear-coverts are light olive ochre, the
sides of the head and neck whitish, washed with olive yellow,
and the throat whitish. The colour of the under parts varies
considerably in different specimens, probably due to sex. In
the one case the general colour is whitish, the crop, which is
furnished with small brown spots, is washed with light olive
brown, the same colour extending down the flanks, and the
under tail-coverts olive buff, marked boldly with brown
longitudinal spots; in the other the colour is whitish, washed
with light olive buff, giving a rich tone which extends down
the flanks: the crop is unspotted.

The under tail-coverts are almost light buff, incon-
spicuously marked with brown longitudinal spots. The tail-
feathers are brown, slightly lighter towards the tip, the
under surface with lavender. There is no downy plumage
in the young.

The plumage of both sexes is in many ways interesting,
since it shows considerable variation. The spots on the
throat are not confined to any one age or sex. The majority
of the males are similar to the one described first, but vary
considerably in intensity of colouring, an absence of spots and
a general richness of tone being apparently complementary
to one another. Between those that are heavily spotted
and those that have the feathers perfectly clear the inter-
mediate grades are numerous. ‘The second one described
simply marks a stage in the deterioration of the plumage
which commences directly coition has taken place and con-
tinues up to the moult. The female described appears to be
the more uncommon form, and I have only seen one other
female that had this peculiarly rich colouring, and hitherto I
have seen no male that could compare with it. The plumage
of the young is also interesting, since an absence of spots in
their case appears to be equivalent to a different tone of
colouring. Whether all the members of a brood develop

t

GRASSHOPPHR-WARBLER

the same colouring, or whether the variations are sexual or
sporadic, further investigation can alone show.

GEOGRAPHICAL DISTRIBUTION.

Owing to certain peculiar characteristics of this species,
naturalists acquainted with its habits are comparatively few,
and consequently our knowledge of its distribution is incom-
plete. We may, therefore, find that certain features in this
distribution, which at the present time appear to us anoma-
lous, may, as our knowledge increases, be partly or possibly
wholly explained.

Great Britain is apparently one of the principal breeding
grounds, although over the whole area it can only be described
as locally distributed. We find it rare in Cornwall, but in the
remainder of the southern counties, including Devon, a regular
but local visitor. Further north it appears to be more
common, and in parts of Oxfordshire, Cambridgeshire,
Gloucestershire, Worcestershire, Shropshire, Yorkshire, Cum-
berland, Durham and Northumberland it is numerous.

The greater part of Wales is unsuited to its habits, con-
sequently it is very local; I have, however, frequently heard
it in Anglesey. Proceeding north, over the border, we find
it gradually becoming scarce; south of an imaginary line
drawn from the mouth of the River Clyde to the Firth of
Forth locally distributed, but north of this line rare, and we
lose sight of it at Arisaig on the mainland and in the north-
west of Skye. In the Orkneys, Shetlands and Western Isles
there is no record of its occurrence. Westward we find it
occurring in the Isle of Man, and then we come to a large
breeding ground in Ireland, where it is generally distributed,
but especially numerous in the counties of Antrim, Dublin,
Wexford and Waterford.

The task of forming a systematic distribution outside the
British Isles is by no means an easy one, and the difficulties
face us directly we cross the Channel.

5)

BRITISH WARBLERS

Commencing with Morocco and Algeria, probably its most
southern breeding range, we find it sparingly distributed.
In Spain we find it occurring in Granada, Malaga, Murcia,
Valencia, and common in the marshes of Santander, also
wintering in the more southern provinces. The records from
Portugal are very scarce, and the bird apparently only visits
the country occasionally : it has been seen near Coimbra.

Crossing the Pyrenees, we find it more generally dis-
tributed and in parts of France common, for instance, in the
provinces of Finistére, Cotes-du-Nord and Morbihan, also
along the banks of the River Var and in Savoie.

Belgium is only rarely visited; it has been found in the
province of Namur. The same applies to Holland to some
extent, but here, perhaps, it is rather more frequent in its
visits and breeds near Haarlem and Cromvoirt: I found no
trace of it in Texel.

Continuing northwards, we find it occurring on migration
in Heligoland, The records, however, become very scanty
as we still continue north. In Holstein it is common, but
rare in Schleswig. It has been obtained on the Island of
Alsen. In Denmark it has apparently only been found during
the autumn migration, specimens having been killed at a
lighthouse on the coast of North-west Jutland, at a lightship
in the North Kattegat and on the Island of Anholt. In
Sweden it has not hitherto been found, but reappears as
a breeding species, though rarely, in the southern parts of
Norway, confining itself principally to the Jideren, Christiania
and Drébak districts.

In Germany it is generally distributed. Working from
west to east, we find.it rare in Hanover, occurring on the
River Marke, and not rare in Baden. The districts near the
Elbe and its tributaries are apparently the most frequented,
especially in Anhalt, Brandenburg, Mecklenburg, near Rostock
and Parchim, and Altenburg. We find it again in Hessen,
and as might be expected, it is partial to the Danube and its
tributaries near Augsburg, Lauingen and Munich, and is not

uncommon in Silesia.
6

GRASSHOPPER-WARBLER

It visits parts of Switzerland, breeding regularly in the
low regions of the basin of the River Aar and the River Thur,
in the districts near Geneva, and in the valleys of Hasli and
Ursern. On migration it is found as high as the Alpine
regions.

Italy is little frequented, and the bird appears to confine
itself principally to the north-western districts, where it is
found in Lombardy, in the Brembana Valley, Piedmont,
Liguria, the neighbourhood of Nizza, and Modena. Little is
known of it in Central and Southern Italy, but it has been
found near Florence in September. From Corsica, Sardinia,
Sicily, Greece, and the Balkan provinces there are no records,

In the Austro-Hungarian Monarchy it is not common,
principally inhabiting the Hansag swamp, and localities in
Pozsony, Feher, Also-Feher and Hunyad. As we get further
east, towards Galicia, it again becomes common, and leaving
Austria, we come to what is apparently its largest breeding
eround—the Russian Empire.

Commencing with Poland, we find it common, especially
in Lublin, then proceeding north, not rare in the Baltic
provinces, the provinces of Pskov and St. Petersburg; still
further north we trace it to the south-east of Lake Onega and
Ustiug Weliki in the province of Vologda It is more
numerous and generally distributed in the central provinces
of Jaroslav, where it breeds on the islands of the Volga and
near Rostov, T'ver, Moscow, Smolensk, and Tula, also in the
district lying between the Rivers Volga and Oka, but from
the middle Volga valley there are no records. In the more
southern parts it is found in the provinces of Kiev, Volhynia,
Tchernigov, also along the banks of the Rivers Bug and
Dnieper. In the Ural district it is commonly distributed
round Bogoslovsk, the tributaries of the River Sosva, and in
the province of Perm; it is also found in the Orenburg
district at the base of the south-west Ural Mountains, in the
plains of the Rivers Sakmara and Ilek, and in the middle
valley of the River Ural. In the Caucasus it is common in

7

BRITISH WARBLERS

the vicinity of Stavropol and the northern slopes, but there
are no records from Trans-Caucasia. Crossing the Caspian
Sea, we find it occurring in the Mangishlak Peninsula,
breeding along the west of the delta of Sir Daria, and in
Western Thian-Shan, visiting Iskandar Kul and the Northern
Pamirs; and still continuing east, we find it on Lake Ala Kul,
and finally lose sight of it on the southern slopes of the Altai
Mountains. There is much to be learnt about the southern
range; at present it has not been found in EKeypt or Tunisia,
but in winter it visits Morocco and Algeria.

Now it must be evident to any one who follows these
details, and at the same time bears in mind certain habits of
this bird referred to in the text, that we have in these habits
a possible explanation of some of the facts which are otherwise
somewhat inexplicable. Since, therefore, it is probable that
at some future date a more intimate acquaintance with these
habits will influence and expand our ultimate knowledge of
the distribution, and in order that the relation they may bear
to the present incomplete state of this knowledge may be
fully realised, I propose to briefly allude to them again.

A partial and in many places almost complete
cessation of the song after pairing has taken place
until the young leave the nest.

A very erratic appearance in any one particular
district from year to year.

Very skulking habits after eight o’clock in the
morning until the evening and during incubation
throughout practically the whole day.

In countries where the bird is seen in the spring and
autumn migration, but is not supposed to breed, especially
where the autumn migration is a comparatively early one,
it is possible that an explanation may be found in the first
of these habits, the bird being overlooked during the quiet
period. But after due allowance is made for these habits,
and the resultant unsatisfactory state of our knowledge, the
geographical distribution remains an interesting one.

8

GRASSHOPPER-WARBLER

It is difficult, for instance, to understand why in Italy
and Hungary—countries affording natural features eminently
suitable to its habits—the bird is scarce. The great plains of
Hungary and the banks of the Danube always appear to me
to be particularly adapted to it, yet I cannot believe that in
a country where the natural life is studied with so great an
interest the bird is overlooked. These remarks apply equally
to the Netherlands, and to some extent Belgium.

The migration across Heligoland, where it occurs not
infrequently in March, April, May, August, September and
October, is completely at variance with the present knowledge
of its distribution. According to Heinrich Gatke, the spring
migration in Heligoland invariably proceeds from west to
east ; but in the case of this species, where? A elance at the
map will show that we are here near 55° N. latitude, above
which, until we reach 25° longitude, the bird can only be
described as rare. ‘Those individuals, therefore, that cross
the island—and the evidence tends to show that their
numbers are considerable—must, after leaving, turn south-
wards into Germany, or continuing east, use this as their route
to the Baltic provinces and Central Russia. But the winter
quarters, as far as we know, are in Morocco, Algeria and the
South of Spain. Is it, then, conceivable that in order to reach
its summer home it would traverse so great and unnecessary
a distance? In the opinion of Heinrich Gatke, the majority
of spring migrants from South Africa on their way to Siberia
do not pass Heligoland, but go direct. It is probable, there-
fore, that there is much to be learnt concerning the northern
breeding range.

LIFE-HISTORY.

Different species vary very considerably in character;
some are lethargic and dull, others are the possessors of great
nervous development, so that the study of some is more likely
to lead to the solution of Nature’s problems than the study of

9

BRITISH WARBLERS

others. To that class in which the different faculties seem
to be specially active belongs the present species, and for
me there is in all their movements, in each action of the
short life they yearly spend amongst us, a fascination
difficult to account for, unless it be in part due to a grow-
ing appreciation of my ignorance of all that they have to
teach me.

Arriving between April 17th and 23rd, they visit us in con-
siderable numbers, yet in a very erratic manner, favouring
one particular district one year and deserting it the next,
neither are they the least influenced in this respect by
climatic conditions ; for instance, in the years 1897 and 1903
they were so plentiful that during the first few hours of
daylight their favourite haunts appeared to be alive with
them, yet these two years were respectively very hot and dry,
and very wet and cold. In Worcestershire, where I have
principally studied this bird, they arrive during the night,
and commence to sing, uninfluenced by the weather, at day-
break; and in the immediate neighbourhood of the spot
they settle in they generally breed, although occasionally one
appears to be a wanderer, singing in a hedge by the roadside,
yet passing on before the next morning. If it were not for
its song, which is penetrating and arrests attention, it would
be a most difficult bird to find, for it is peculiarly skulking
in its habits, especially after eight o’clock in the morning until
about four in the afternoon. It spends the greater part of its
time either on the ground in the dense undergrowth, creeping
in low thick bushes a few feet high, or searching tall thick
hazels and hedgerows for food ten or twelve feet from the
eround, but never appears to frequent trees; and has at all
times, even when unconscious of any human presence, a.
ereat aversion to showing itself in the open. During incuba-
tion, when the song partly ceases, these skulking habits are
more marked, since both sexes go to considerable trouble to
conceal themselves.

They choose for themselves different and very varied spots,

10

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GRASSHOPPER-WARBLER

but perhaps those most frequented are osier-beds, which a
year or so previously have been cut down; small coppices
with plenty of thick undergrowth; spaces in large woods
where timber has lately been felled, and where consequently
scrubby bushes and luxuriant vegetation abound; waste land
covered with gorse; swampy, but not too wet, ground with an
abundance of Juncus effusus; clover fields and thick hedge-
rows either by the roadside or next to fields of corn; thick
undergrowth of some description seems, however, to bea
necessity.

On their arrival the males can easily be detected by their
song—for applying the word “song” to the vocal efforts of
my friends I must apologise, I know not what other word
describes it—which when once heard can never be forgotten,
and from which they have earned the name “ Grasshopper ”’ ;
yet when with bill wide open and body quivering they utter
their monotonous “reel,” there is little resemblance to the
insect. A more minute knowledge of their ways is required to
enable one to appreciate whence the name originated. They
are by no means difficult birds to approach, and with ordinary
precaution it is possible to get concealed quite close to them,
and to hear the low whisperings from which they take their
name. Let us suppose for a moment we are concealed within
a few yards of one of them. There, on a branch low down
in some nut-bush, he sits wrapped in meditation; now he
commences to sing, and the power thus expended is apparent
from the vibration, which is becoming almost painful to the
ears, when he stops, and in turn in the far distance we hear
others answering, first on one side, then on another. But
watch him closely ; with throat still distended and bill slightly
open, he turns his head from side to side, and the sounds
from our imaginary birds in the distance coincide with the
turning of his head; thus the truth dawns upon us. He is
certainly a wonderful ventriloquist, and though perhaps not
conscious of his powers, yet he is most deceptive, and I
have often been puzzled when trying to locate his position,

set:

BRITISH WARBLERS

especially when he will do nothing but whisper, for this
ventriloquistic effect is not so marked when singing loudly.
Until pairing has taken place the song is almost incessant,
morning and evening, rare during the daytime, more frequent
at night; afterwards, and until the young are able to take
care of themselves, it almost ceases, until it becomes a mere
apology of the song of the mating season, often reduced to
a few crackling notes, heard on a hot afternoon or during
the first hour or so of dawn. To this partial cessation of
the song may possibly be traced the belief that in certain
districts the birds do not stop to breed, but are only to be
found on their way to their breeding quarters; yet it is
characteristic of many of the warblers, but perhaps more
marked in the one under consideration; and thus the theory
of the male lightening the female’s task of incubation by
song becomes a fanciful one, having little foundation of fact
to recommend it, and against it the weighty evidence of
Nature who always strives for a more complete concealment
of her children. Before the arrival of the females, the males
during the first few hours of daylight are very fond of playing
with one another; and although their games are not to us
very interesting, yet considerable energy is expended on them
by the birds themselves. One male darts off after another
one, pursuing him with very rapid flight into some thick
bush; here he chases him up and down and along the
branches with tail outspread and wings extended and slowly
flapping, his behaviour being exactly similar to that during
courtship, occasionally making use of a very curious note,
much like the scolding of the Garden Warbler (Sylvia
hortensis) when its nest ig approached; and this sound
appears to be produced with considerable exertion.

On the arrival of the females, about April 27th, ten days or
so after the males, the courtship is immediately commenced.
No period in the life-history of the individuals of any species
is so interesting to watch or so full of significance as this.
Every nerve is strained to a degree which makes incidents in

12

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GRASSHOPPER-WARBLER

their ordinary lives appear almost commonplace. Listening
during these few days at daybreak to the song of the males,
it is quite possible to tell whether any females have arrived
during the night; at such times it is intermittent, but very
vigorous while it lasts—a pause, then a short trill, a longer
pause and a longer trill, and so on, according to the time
the male is able to tear himself away from the attractions
of the female. She, meanwhile, walks or runs—as a rule
it is more of a sedate walk—amongst the dense under-
crowth, at times threading her way through the branches
of some small bush, occasionally pecking, or pretending
to peck, at something as she passes, with an air of com-
plete indifference to the attractions of her lover, never even
pausing to look behind. He, however, sometimes leaves
her either to sing, or with quick darting flight to chase
away another male, who, in his opinion, has approached too
closely, or for a more lengthened period, possibly to annoy
other males engaged in the same task as himself. But she
is immediately aware of his desertion, and if too long away
becomes impatient, stops her aimless wandering in the under-
erowth, flies to a low branch in some bush, assumes an air
of anxiety, and commences to call vigorously till he returns,
using as her call-note a single “tic,” sometimes the “ tic tic
tac.” ‘The positions assumed by the male when following her
—sometimes he walks, sometimes runs along the branches or
on the ground—are very beautiful. ‘The tail is spread out and
either raised or lowered, the wings fully extended and slowly
flapping up and down, the feathers on the back and head
are raised, those on the body thrown out, and the throat dis-
tended. The effect as he walks along some horizontal branch
is very striking, and is enhanced by the curious habit, common
amongst so many species, of carrying something in his bill,
a piece of dead grass perhaps, but more frequently a dead
leaf, and when two or three of the latter, stuck together, are
picked up and carried for some distance the effect is striking
and at first somewhat startling, giving one the impression

13

BRITISH WARBLERS

in the dim morning light of a stoat-like animal crawling
along the branches. This habit, whatever the origin, seems
now to be an indispensable part of the courtship and is
very strongly implanted in the male; for, before following
the female, the leaf or piece of grass is searched for with
impatience, as the female does not stop but calmly pursues
her way, the result being that whatever he first gets hold of,
and can conveniently carry, is sufficient for his purpose.

In referring to this period of the bird’s life, I have used
the simple term “cowrtship,” instead of the usual display
followed by selection. Fascinated by a picturesque theory,
I have, previous to this, attempted to interpret facts accord-
ingly instead of being ad utrumque paratus, but closer study,
devoted for some years to this courtship, convinces me that
sexual selection as a rational explanation of the phenomena is
impossible: the following reasons have been mainly instru-
mental in forcing this conclusion upon me.

Selection is impossible unless there are two or
more males to be selected from.

I can find no evidence of this being a general rule.in
Nature, for although a second male occasionally approaches -
the zone of courtship and is consequently chased away by the
first male, yet he is in no sense a rival, making no attempt to
perform in the same extravagant manner; and if these two
males were in competition for the female, and if such com-
petition were to be decided solely by her choice, then, not only
ought they both to display their plumage, but, unless the
female is to be credited with sufficient mental power to enable
her to carry, for her further consideration, the varying
characteristics of each male in her mind, such display ought to
take place simultaneously.

“Tf the male assumes these attitudes in order that
the female may see him to the best advantage and thus
be enabled to make her choice, then she ought to be
watching him when thus occupied.”

This is by no means the case, for she is generally walking,

14

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ATTITUDE DURING COURTSHIP.

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GRASSHOPPER-WARBLER

running, or flying away from her suitor—indeed, I cannot call
to mind a single instance in which I have, even for a brief
moment, seen the female looking at the male while assuming
these positions—and therefore has her back turned to him
when we ought to expect her to be examining the details of
his plumage. It might be here argued that for the female to
get the general effect of the male’s display would, through
such immense periods of time, be sufficient to account for the
beauty of the plumage; but I cannot bring myself to believe,
even if the other and more weighty objections could be
removed, that such a casual inspection could be the means of
developing minute and beautiful patterns, the very delicacy of
which must surely cause us all to marvel. This, however, we
know, that as a bird grows older the colours of the plumage
become intensified, and even the plumage itself becomes finer
in quality, and amongst the Warblers this difference is go
marked as to at once arrest attention. He, then, who
supports the principle of selection must admit that the older
males being more richly coloured would be first selected; he
must also admit that the offspring of these more successful
males could inherit no advantage, inasmuch as the success of
their parents was due solely to laws of continuous growth.

“Karly in July a courtship, similar in every
respect to the one in spring, occurs amongst the
Sedge- Warblers.”

Admitting the principle of selection, we must nolens volens
assume this to be a selection taking place before the second
brood, conclusively proving the choice of the female in the
first instance to have been a mistake.

“The ultimate production of the most healthy and
most beautiful offspring by the selection of certain
males is, without a corresponding selection amongst the
females, impossible, and of the existence of such selec-
tion in any form there is no evidence.”

Individuals of both sexes vary very considerably both in
intensity of colouring and actual quality of plumage. -As an

15

BRITISH WARBLERS

instance, the females of the present species are occasionally as
rich in colouring as the finest males. Here, then, the im-
potency of sexwal selection becomes apparent, for unless the
females that choose the more beautiful males are themselves
the more beautiful of the females, they will neutralise the
effect of their own selection.

Although this is a truism which appears to me to be
difficult to controvert, yet I must admit that we are here
confronted with two difficulties, one an incomplete knowledge
of the laws relating to heredity, the other an ignorance of the
influence exerted by the female upon her embryo, which is
profound. We do not even know whether heredity and this
influence are distinct, or inseparable, merging into one another
by very gradual stages. Yet this influence is a potent factor
in all life, including man.’

Little importance need, however, be attached to these
difficulties here, since they depend upon the supposition that
characters acquired by one sex can be transmitted to that sex
only. For instance, if the dull-coloured female Grasshopper-
Warblers in a given area were to pair with the bright-coloured
males, and the bright-coloured females with the dull-coloured
males, the result in time would not be the continuation of these
same conditions, but the gradual annihilation of the bright or
the dull colours. It is inconceivable and contrary to the facts
in Nature to suppose that the colouring of the female offspring
could be in no degree influenced by the male parent, and
vice versa.

It has been suggested that the more vigorous females
would be the first to breed, but this is a supposition made
only to escape a difficulty, evidence in support of it being
completely lacking: and since these same females vary
individually very considerably in their colouring, we should
at once, providing evidence were forthcoming in proof of

1 Tt ig indeed difficult to understand the callousness with which this
knowledge is treated in human life at the present day.

16

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_ such a supposition, be led to assume that colour was solely
dependent upon and inseparable from vigour; thus the
necessity for calling in such remote possibilities as esthetic
sense and selection in explanation of the phenomenon would
be removed.

Let us compare the courtship of the famous Argus
Pheasant with that of any of the dull-coloured Warblers, for
instance, Savi’s Warbler (Locustella luscinioides). Here we
have two cases, extreme as regards development of colour and
shading, yet analogous in all other respects. The Argus
Pheasant, when in presence of the female, spreads out and
raises its tail and wings, the ocelli on the latter being thereby
fully displayed; Savi’s Warbler also under similar circum-
stances spreads out his tail and wings, the latter being
very slowly waved up and down. These actions are identical,
and evidently spring from the same cause, but no one can
say that Savi’s Warbler, in thus performing, has any special
beauty to display. If, then, we say that the Areus Pheasant
is conscious of the ocelli, of what is Savi’s Warbler conscious ?
Clearly we are no more justified in imputing consciousness
to the actions in the one case any more than we are in
those of the other. :

‘The courtship of birds presents many curious
features; none more so than the spreading out of
the wings and tail, commonly known as display, but
these actions are not confined solely to courtship.

I have described the males of the species under considera-
tion, both when playing with one another prior to the arrival
of the females, and when feeding their fully fledged young,
as behaving in an exactly similar manner. The male Chiff-
chaff, before the arrival of the female sometimes seems to be
seized with a sudden ecstasy. He spreads his tail and jerks
his wings, singing to himself quickly and quietly; also when
a hawk approaches his breeding-quarters too closely he will
fly into the air, slowly flapping his wings, and hurriedly singing
in a similar manner to his courtship. I will give one other

17

BRITISH WARBLERS

instance: he male Blackcaps, when excited with one another,
or with another species, such as Sylvia hortensis, assume the
same positions as when courting the females. Tio what do
these facts point? Certainly not to a conscious display.

Further evidence, details of which I need not enter into
here, prove that. all these actions are intimately associated
with the development and stimulation of the sexual organs.

When we come to consider all these facts, when, on the
one hand, we find evidence of so strong a character opposed
to the theory of seawual selection, and on the other, find that
actions similar in every respect are frequently performed at
other periods of great excitement in a bird’s life, can we doubt
the probability of their being solely reflex in kind?! I think
not. All the difficulties would thus be removed, the second
courtship of the Sedge-Warbler—a courtship which, although
I have not hitherto seen it, yet in my opinion undoubtedly
occurs amongst other species—is explained; and there would
only remain the general argument, namely, that it is impossible
to believe that all the trouble which many species apparently
take with their courtship is purposeless: and this argument
may be raised against the theory of reflex action; but because
I exclude conscious choice from being in any way directly
responsible for these actions, I by no means wish to imply
that they are purposeless, neither am I prepared to enter
into further explanation here, for I hold that with the very
meagre knowledge that we, even at the present day, possess
of all the facts relating to this most interesting subject, it
would be unwise to attempt to formulate a theory in explana-
tion of their ultimate purpose.

The peculiar manner in which these birds walk along

‘In his ‘‘ Last Words on Evolution,’’ Professor Ernst Haeckel considers
“sexual selection” of the greatest importance, both for the general theory
of evolution and also for psychology, anthropology, and esthetics. Since
the importance of it is so great, no labour, surely, ought to have been spared
to have verified the completeness of the fundamental evidence upon which
it was originally based ?

18

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GRASSHOPPER-WARBLER

horizontal branches of very small dimensions is really very
graceful; one wonders how they keep their balance and
yet preserve such a very aristocratic appearance. Climbing
in this way up some bush—crawling, perhaps, is a_ better
description—their whole appearance is peculiarly mouse-like,
especially as wood-mice are themselves very fond of climbing
up and sitting in small bushes, such as hazel, elder, &c.

The nest, composed principally of dried grass interwoven
with dead leaves and bracken, is, even when the young are
hatched, most difficult to find. Placed sometimes almost on
the ground, sometimes a few feet above it, it may be found in
very different situations, such as in the centre of a clump of
Juncus effusus, underneath a thick mass of the same rush
overhanging a small watercourse, in the middle of clumps of
long dead grass, and low down in thick gorse bushes. During
incubation, which lasts about sixteen days, the female seldom
leaves the nest. I once, in June, saw a pair playing with
one another as it was getting light, between two and three
o’clock in the morning; they were chasing one another, and
the female settled and rested awhile on a branch quite close
tome. When disturbed on her nest she will slip off quietly
on to the ground and be quickly concealed, but if not too much
alarmed will return, threading her way through the grass in
the same stealthy manner. If lucky enough to find a nest
with young, watch closely the parent birds; time thus spent
will by no means be wasted. One nest, I remember, placed
low down in a gorse bush, and in a great measure hidden
by long grass, gave me an exceptional opportunity of doing
this. The locality I found by luck, for, when walking close
by, one of the parent birds was disturbed and flew out, but
it was only with considerable difficulty that I found the
actual nest. Crawling under the gorse and lying flat was
useless, as one or other of the birds would walk within a few
feet of my face and stare; to stand up seemed hopeless, but
nevertheless it ultimately succeeded, and while I was
remaining perfectly still, patiently enduring torments caused

19

BRITISH WARBLERS

by insects, one of them, with a billful of food, went to the
nest within a few feet of me. Upon my taking up the same
position at daylight, the birds, losing a great deal of their
fear, continued with the care of their young. In going to
or leaving their nest they always ran, and so much do they
resemble mice that I was once completely taken in, believing
that what I saw was in reality a mouse. One of the parent
birds once raised its wings in a threatening attitude, but this
was the only occasion on which they showed any sign of
objection to my presence. A note was frequently uttered by
both sexes, but it seemed to me to be only a call-note to the
young, probably to keep them quiet: the male sang at
intervals very quietly, and the young, when being fed, uttered
tiny squeaks.

When feeding the young they bring billfuls of insects, or
to be more accurate, lumps of squashed insects, and by
examining these lumps it is possible to find out what their
food consists of. Wait until one approaches you on the way
to the nest, then make a sudden movement or a step forward,
and the food will in most cases be dropped. Their dexterity
in collecting the insects is marvellous; even when their bill
appears to be full they still continue darting at an aphis
here and a gnat there, adding them to the lump, yet never
dropping any of that already gathered, so quick are their
movements.

The young leave the nest when only a few days old, and
until able to fly are most difficult to find, concealed as they
are in the dense undergrowth; but in the study of Nature
it may with considerable truth be said that “all things come
to him who waits.” Having ascertained the position of a
nest from which the young have only lately gone, go there
soon after sunrise and get well hidden. You will hear the
hum of insects commencing and gaining in strength, as in
response to the intensifying rays of the sun each one begins
his daily toil; you will hear an insect with a very small
far-away squeak, which for a time will puzzle you, but since

20

WARBLER

CR we te ile HE. maNae

GRASSHOPPER-WARBLER

it continues at frequent intervals to attract your attention
it will make you wonder whether it may not after all be some-
thing more than an insect, and your suspicion is well founded.
It was, indeed, some time before it dawned upon me that the
owner of the voice might be a young Grasshopper- Warbler,
for no tinier sound could come from the throat of any bird
out of the nest. And now how to find him? Often have I
searched in vain; in gorse it is well-nigh hopeless and best
avoided, in osier-beds and waste swampy ground patience only
is required to crown one’s efforts with success.

Crawling amongst the long tangled undergrowth into the
middle of hazel bushes, where the foliage is so dense that light
can scarcely penetrate, | attempt to follow the sound. Hach
time on hearing the squeak I crawl a little closer until quite
certain that the little bird is within a few inches of my face,
but quite uncertain whether to the right or left or straight
in front of me; still, by waiting, and with head inclined first
one way then another, noting the direction the sound appears
to come from, I make up my mind as to the exact spot,
divide the undergrowth with the greatest care, and am
amazed to hear the squeak come from exactly behind me.
Ii this is not ventriloquism I know not what else to call it;
it is an even more effective deception than the adult birds
are capable of. At this age the young have very few feathers,
and their naturally long legs give them a rather clumsy
appearance; but in reality they are capable of running quite
fast, and to this fact, until an opportunity occurred of
studying them more closely, I was inclined to partly attribute
their ventriloquism ; for when unconscious of being watched,
they stand huddled up on the ground, their heads drawn close
into their bodies, wings drooping, feathers loose and eyes half
closed, and off and on they raise their heads for a moment,
open their beaks and utter their little squeak, turning their
head in expectation of food first one way then another,
apparently unconscious of the ventriloquistic effect produced.
The parent birds sometimes show very little solicitude for

a1

BRITISH WARBLERS

their offspring, even under the most trying circumstances ;
for when I have been sitting with a young one on my hand
they silently came and peeped through the branches, dis-
appeared for a moment, and looked again from the opposite
direction, but made no sound, neither did they attempt in any
way to entice me away. At other times when, attracted by
the call, I have attempted to find the young, the female has
shown considerable anxiety. She would utter a note which
was a peremptory signal for the young to keep quiet, neither
would any one of them, although scattered and a considerable
distance apart, call again until she gave another signal;
waiting for which signal considerable patience on my part
was required, for the slightest movement would make her
suspicious, hidden though she appeared to be, and when her
suspicions were aroused she would immediately come up to
the top of the gorse bush on which she was and watch my
movements closely, nor were her suspicions allayed until I
had kept quiet for a considerable time. She seems herself
to do all the feeding of the young after they have left the
nest, the males leading a lazy life, singing and playing with
one another. One instance, and this a curious one, I have
seen of the male feeding the young when almost full grown.
It was in a large field of clover, and I could find no sign of
the female at all. The young were scattered considerably,
and at intervals between the feeding the male sat on a post
and sang. When actually feeding one of the young he would
spread out his tail and wings and slowly wave them up and
down. ‘The young, when able to fly a little, occasionally come
out of their hiding, but if pursued and made to fly they very
soon get tired, and if driven into an open piece of ground,
where there is no cover for them to conceal themselves
in, they become completely lost. This I have proved by
manceuvring them on to patches of burnt gorse: here they
would run aimlessly about and make no attempt to conceal
themselves. |

I have once seen, on June 17th, two of the adult birds

122

GRASSHOPPER-WARBLER

apparently on the point of pairing. The male was excitedly
following the female, singing at intervals, and at times
bowing slightly ; but this is the only evidence I have that two
broods may be reared in the same season, while on the other
hand the general behaviour of the males, when the young
are fully fledged, is very different to this, for they sing for
about a fortnight, that is to say, during the latter part of
June and the beginning of July, then become restless and
wander away from their breeding quarters. During these
last days of June, if the weather is warm, their song can be
heard to advantage in the evening and on through the night,
rising and falling as the head is inclined first this way and
then that. It is surprising how the song will penetrate, for
it can be heard without difficulty a quarter of a mile away. It
often happens, too, at this time of year, that the only other
sounds in the night-time are the “churring” of the Nightjar,
the craking of the Landrail, and the hissing of the young
Brown Owls, sounds which are well in keeping and only
tend to emphasise the stillness of the night.

Although their song is so peculiar, yet there are a few
notes in the song of the T'ree-Pipit and Wren for which they
may easily be mistaken, and often when listening at day-
break for these Warblers I have been completely deceived
by these few notes, which, at a distance, with the remainder
of the song lost, are barely distinguishable from those of the
Wren or Tree-Pipit. The range of call-notes they use seems
to be small; the usual one is something like that of the Robin
or Blackcap, a “tic tac tac,” and this note seems to be used
principally when alarmed. The note which the female uses
to call the male, or to put the young at their ease, is usually
a single “¢ic”’; on the other hand, this single “tic” quickly
and loudly uttered may also be one of alarm; it seems, there-
fore, that it depends more on the manner in which the note
is given than on the actual note itself. There is no difference
in the note of the two sexes.

Climatic conditions seem to influence them to a certain

23

BRITISH WARBLERS -

extent. In very wet weather they mope, and when in this
frame of mind it is possible to approach them very closely and
watch the vigorous attempts they make to dry themselves.
There is also at certain times towards the end of June a
cessation of their song for a few days, which, owing to its
curious spontaneity, must, I think, be due to changes of
weather. In cold frosty springs I have never been lucky
enough to find sufficient nests to enable me to form an
opinion as to the effect of frost on the eges, so disastrous
to those of many species.

These birds are entirely insectivorous, aphides in particular
forming a great part of their diet; wood-lice, spiders, small
moths belonging to the family Geometre are also taken; I
have also seen the males in June feeding upon the larve of
Chimatobra brumata.

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